Madagascar, with descriptions of eight new species and

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1 Some Phytoseiidae (Acarina: Mesostigmata) from Madagascar, with descriptions of eight new species and notes on their biology by LeoBlommers Laboratory ofexperimental Entomology, University ofamsterdam, The Netherlands ') Abstract (Nietner, 1861). Most of these results were negative. Only, Typhlodromus contiguus, Amblyseius tulearensis and A. sundi The taxonomic portion of this paper discusses 27 species of mites of the family Phytoseiidae known to occur on plants in Madagascar. Eight new species are described: Phytoseius (Phytoseius) can regulate population numbers of the first prey species under circumstances of low webbing and/or additional food. Iphiseius degenerans, and possibly Amblyseius can rotundus, control Oligonychus coffeae in their area of distribution. betsiboka. Ph. (Pennaseius) onilahy, Amblyseius (Amblyseius) hova, A. (A.) ankaratrae, A. (A.) boina, A. (A.) sakalava, A. (A.) trichophilus and A. (Proprioseiopsis) tulearensis. INTRODUCTION The following species are recorded for the first time from the island: Typhlodromus (Typhloseiopsis) contiguus Chant, 1959, Phyloseius (Phytoseius) crinitus Swirski & Shechter, 1961, Ph. (Ph.) intermedius Evans & Macfarlane, 1961, Ph. (Pennaseius) hongkongensis Swirski & Shechter, 1961, Ph. (P.) amba Pritchard & Baker, 1962, Iphiseius (Iphiseius) degenerans (Berlese, 1889), I. (Trochoseius) gongylus Pritchard & Baker, 1962, Paraphytoseius multidentatus Swirski & Shechter, 1961, Amblyseius (Amblyseius) hima Pritchard & Baker, 1962, A. (A.) deleoni Muma & Denmark, 1970, A. (Proprioseiopsis) sundi Pritchard & Baker, 1962 and A. (P.)peltatus Van der Merwe, Additional descriptions are given for most of these species. The male of P. multidentatusis described for the first Phytoseiid mites rank among the most important natural enemies of phytophagous mites and therefore have received increasing attention during the last decennia. The number of described species has grown from less than 20 in 1951 (Nesbitt), to around 1000 at the moment. This paper is the last in a series comprising an inventory of phytoseiid mites as predators of spider mites (Tetranychidae) conducted between 1970 and 1973in Madagascar. time. New distribution records are presented for the following species: Typhlodromus scytinus Chazeau, 1970, the T. (Anthoseius) gutierrezi-chazeaui group, Amblyseius (Amblyseius) SYSTEMATICS masiaka Blommers & Chazeau, 1974, A. (A.) bibens Blommers, 1973, A. (A.) rotundus Blommers, 1973, A. (A.) brevipes Material and methods Blommers, 1973 and A. (A.) tamatavensis Blommers, The simultaneous occurrence of spider mites and other small arthropods on the plants on which the phytoseiids were collected is noted. A final chapter summarizes the results of experiments devised to see whether the collected phytoseiid mites can be effectively predators of spider mites, i.e. of Tetranychus neocaledonicus André, 1933 and sometimes Oligonychuscoffeae All phytoseiid mites were collected on plants by R. Blommers-Schlosser and the present author unless stated otherwise. The specimens were preserved in 70 per cent alcohol, cleared in 50 cent lactic acid and mounted in Marc per Andre's medium. As the knowledge of natural relationships ') Author's address: present Laboratory of Parasitology, University of Leiden, Rapenburg 33, Leiden, The Netherlands. within the Phytoseiidae is still inconclusive, I have chosen a rather conservative generic and subgeneric division, which agrees in general with

2 Two Two Four Our BIJDRAGENTOT - DE DIERKUNDE, 46 (1) the one used by Van der Merwe (1968). The following are exceptions. According to its revised definition (Muma & Denmark, 1968), the subgenus Proprioseiopsis Muma, 1961, of Amblyseius 21.V at Ifaty, in sea dunes 10 km N. of Tul6ar. T. gutierrezi and T. chazeaui are intimately related, and appear to form of part a more extensive species group. Both species were described from the region of Tulear, where it is easy to distinguish them. should contain also those species in which dorsal T. gutierrezi seems to be restricted to the more setae J2 are absent, and Phytoseius Ribaga, 1904, humid biotopes in the direct neighbourhood of and Paraphytoseius Swirski & Shechter, 1961, should be raised to generic rank. Tulear city, while T. chazeaui is a form of the sea dunes with sclerophyllous vegetation. The nomenclature of the dorsal setae is that Our material from Majunga province, proposed by Lindquist & Evans (1965) and however, contains forms which we are unable to adapted for Phytoseiidae by Athias-Henriot (1966) Blommers & (cf. Chazeau (1974) and fig. Dimensions are 1). given in micrometers. The identify for the moment. in various They show, degrees of mixing, features which are not only to proper both aforementioned species but also author's collection numbers accompany the col- to South African T. paganus Van der Merwe, lection data. 1968, and T. vescus Van der Merwe, More Spider mites collected concurrently with the material is needed to understand the species phytoseiid mites have been identified by Dr. J. complex. Gutierrez, and are mentioned eventually together with other small arthropods. The entire collection of Malagasy Phytoseiidae is deposited in the Institute of Taxonomic Zoology (Zoologisch Museum) ofthe University of Amsterdam. The following specimens belong to this complex: 1 9 and <5 (M I series) on a species of Asclepiadaceae, (M 3 series) and (M 14 series) on unknown plants, 1 9 and 1 (j (M7 series) on Rhopalocarpus lucidus (Rhopalocarpaceae), (M 9 series) on Bauduinia sp. (Caesalpinaceae), 1 <} (M 13.1) on Flacourtia ramontschi (Flacourtiaceae), and 2,5 (5 (M 16 series) on Diospyros sp. (Ebenaceae), all from sand Descriptions dunes near Katsepy (Majunga), 26.IV Further, 3 99 (M 6 series) on Ricinus communis (Euphorbiaceae), 1 9 (M 15.9) on Leonotis nepetaefolia (Labiatae), (M 17 series) Typhlodromus scytinus Chazeau, 1970 on Carica papaya (Caricaceae) at Ampijoroa (Majunga province), 24. IV In the same locality, 599 and 2 33 Typhlodromus scytinus Chazeau, 1970: 3, figs. 17. $9 (T 18 series) collected on grapevine (Vitis vinifera), 1. III at Tananarive-Ambodrona. (M 33 series) also on R. communis and 1,5 (M 34.4) on Leptadenia madagascariensis (Asclepiadaceae), 30.XI One,5 (M 12.5) on Pueraria javanica (Papilionaceae) at Ambato Boeni, 24.IV.1972; in the same place, and 4 (j (M 26 series) on Alchornea sp. (Euphorbiaceae), Typhlodromus (Anthoseius) gutierrezi Blommers, IX Typhlodromus (Typhloseiopsis) contiguus Typhlodromus (Anthoseius) gutierrezi Blommers, 1973: 109, figs. 17. Chant, 1959 Four 9 9 and one <} (64 series) from Euphorbia hirta, 9.III.1971; one 9 (63.3) from Abelmoschus esculentus (Malvaceae) and two 9 9 (54 series) from Corchorus trilocularis C. (Tiliaceae), both ; one 9 (65.10) also from trilocularis, 2.1 V. 1971; all these specimens collected in the city oftul6ar. Typhlodromus (Typhlodromus) contiguus Chant, 1959a: 29, figs. 1 6; Chant, 1959b: 50, figs Typhlodromus (Typhloseiopsis) contiguus; Ehara, 1967: 71, figs and four <j $ (M 223k series) from a rearing starting with specimens found on several plants (Phaseolus sp Papilionaceae; Leonotis sp., Labiatae and Typhlodromus (Anthoseius) chazeaui Blommers, 1973 Ipomoea sp., Convolvulaceae) near homestead gardens in coastal forest, a few miles S. of Katsepy (Bombetoka Bay, opposite Majunga), 27.IV Typhlodromus (Anthoseius) chazeaui Blommers, 1973: 110, figs (B 39 series) from Solatium sp., Diagnosis. specimens agree well with the available descriptions of this highly characteristic Typhlodromus species. Some dimensions in the female are slightly different: length dorsal shield 310, jl 25, j3 45, Z1 89.Z4 96, Z5 160, s3 70,s4 80,

3 Ten Six The The This PHYTOSEIIDAE The 82 L. BLOMMERS - FROM MADAGASCAR VL1 71, macrosetae tibia and basitarsus genu, IV rearing started with specimens collected on Urena lobata of the same length: about 60. (Malvaceae) along Route Nationale 4, km 491, near Amboromalandy, 11.IX Three paratypes (2 9 9 ar>d 1 <$\ M 28 series) from Combretum villosum (Combretaceae) on the Phytoseius (Phytoseius) crinitus Swirski & Shechter, 1961 Phytoseius (Dubininellus) crinitus Swirski & Shechter, 1961: 102, figs. 810 (9); Swirski & Amitai, 1966: 11, figs. 13 (cj); Denmark, 1966: 66, fig. 27. Phytoseius (Phytoseius) crinitus; Ehara & Lee, 1971: 71, figs (series T 19) were collected from grapevine ( Vitis vinifera; Vitaceae) at Soavina, Miantso, Tananarive province, altitude about 1300 m, on 8.II.1972; one 9 (T 10A.4) from lemon (Citrus limon; Rutaceae) in Tananarive-Nanisana, altitude 1250 m, 21.1X Two 9 9 (A 23.3 and A 42.3) were collected from Psidium guayava (Myrtaceae), the first at on Ivoloina, Tamatave, 25.V , the second in the city of Tamatave, 5.VIII same spot and the same date as the holotype. Differential diagnosis. of the pattern dorsal setae, R1 being absent, places this species within the subgenus Phytoseius Ribaga, Only a few species within this genus possess a single pair of pre-anal setae on the ventri-anal shield. P. betsiboka differs from them in the extreme length of seta s4 which is considerably longer than any other dorsal seta. The presence of a large notocephalic pore caudad of seta z5 is unique among known species of this subgenus, but is found in a great majority of species in the subgenus Pennaseius Pritchard & Baker, Discussion. specimens from Soavina and Ivoloina agree exceptionally well with the original description of P. crinitus. A approaches Phytoseius ferox Pritchard & Baker, 1962, in having seta s3 shorter and macroseta genu IV longer; dorsal setae s4 and Z4 are not divided in this specimen. Description. Female: Dorsal shield smooth, 280 long and 140 wide; incised near seta r2; with at least ten pairs of pores, two of them large and surrounded by a distinct rim; fifteen pairs of setae on the shield; the longer thickened, with of rows spines along their length; their length: jl 30, j3 57, Z1 86, Z4 98, Z5 114, s3 23, s4 166, r2 54; the remaining setae (j4, j5, j6, J5, z4, z5, s2) smooth and minute, Phytoseius (Phytoseius) intermedius Evans & Macfarlane, 1961 less than 7 long. Peritremes reach nearly in front of setae j 1. Phytoseius (Dubininellus) intermedius Evans & Macfarlane, 1961: 587, figs. 13; Denmark, 1966: 70, fig. 29. Phytoseius (Phytoseius) intermedius; Ehara, 1972: 170, figs Phytoseius (Phytoseius) yira Pritchard & Baker, 1962: 227, figs Sternal shield not well visible, poorly sclerotized. Genital shield as usual, 75 wide. Ventri-anal shield 93 long and 46 wide, without pores and with a single pair of pre-anals. Surrounding membrane with six pairs of pores, two pairs of metapodal platelets, and five pairs of setae; VL1 99 (A 26 series) from guava (Psidium thickened and serrate, 78 long. guayava; Myrtaceae) in Tamatave City, 28.VII Leg IV with four blunt macrosetae: on both genu and tibia 18 long, on basitarsus 40, distitar- Diagnosis. Malagasy specimens agree sus 42. Some other setae might be blunt too. well with the descriptions given. of this species already Remaining legs without distinct macrosetae. Length tarsus IV (including basitarsus) 196. Both digits of chelicera about 23 long. Fixed Field note. phytoseiid was accompanied by a not yet described spider mite of the genus Oligonychus Berlese, with digit four teeth, arranged as figured; movable digit with one tooth. Spermatheca as figured. Major duct 29 long; cervix containing spermatophore 46 long. Phytoseius (Phytoseius) betsiboka sp. n. (figs. 16) Male: Dorsal shield 240 long and 130 wide. Arrangement of pores as in the female. Length of the large serrate setae: jl 23, j3 48, Z1 59, Z4 68, Holotype 9 (serial no. M 30k.2) and seven paratypes (M 30k series: 3 and 4 $9 tfcj) from a mass Z5 61, s3 21, s4 132, r2 40. Remaining setae minute.

4 BIJDRAGEN TOT DE DIERKUNDE, 46 (1) Figs. 16. Phytoseius betsiboka sp. n.: 1, dorsum ; 2, chelicera ; 3, spermatheca ; 4, leg IV ; 5, genital and ventri-anal shield ; 6, chelicera. Figs Phytoseius onilahy sp. n.: 7, leg IV ; 8, spermatheca with four spermatophores; 9, mediolateral part of dorsal shield.

5 Holotype The A 84 L. - BLOMMERS PHYTOSEIIDAE FROM MADAGASCAR Ventri-anal shield as usual, 100 long, probably separate from the peritremal shields, with three pairs of pre-anals and no pores. Four blunt macrosetae on leg IV; on basitarsus 34 long. Length of tarsus IV 152. Chelicera as figured; both digits about 16 long. Fixed digit with three teeth. Movable digit with one; spermatophoral process gibbet-shaped, major portion 11 long, branch 5. wide, without with one pores, pair of pre-anal setae. Surrounding membrane with setae and pores arranged as in P. betsiboka and two pairs of metapodal platelets; VL1 95 long. Leg IV with four blunt macrosetae; on both genu and tibia 20, both on basitarsus and distitarsus 36. Some other setae on leg IV are also blunt. Remaining legs without distinct macrosetae. Length tarsus IV (including basitarsus) 179. Both digits of chelicera about 24 long. Number Field note. and of teeth as in P. arrangement betsiboka. Eotetranychus paracybelus Gutierrez, 1967, was found once together with this species (M 30). Spermatheca as in P. betsiboka; major long, cervix 30 x 11. duct 27 Etymology. This species is named after the largest river of Madagascar which joins the sea near Majunga. Male: Dorsal shield 223 long and 134 wide. Arrangement of pores as in the female. Length of setae: jl 25, j3 55, Z1 59, Z4 68, Z5 66, s3 23, s4 116, r2 45, R1 10. Remaining dorsal setae minute. Ventri-analshield as in P. betsiboka, 96 long. Phytoseius (Pennaseius) onilahy sp. n. (figs. 79) Four blunt macrosetae on leg IV; on basitarsus 34, on distitarsus 30. Length oftarsus IV 143. Chelicera not easily discernable, but probably 9 (serial no. 47.4) and 12 paratypes similar to P. betsiboka. (series 47 and B 13; and 5 #<J) were collected from Sida sp. (Malvaceae) in the garden of the Agricultural Station Remark. single dorsal seta J2 is present in intulear,9and24.iii.i971. two specimens: and 47.7 <J. Differential diagnosis. possession of setae Discussion. R1 places P. onilahy into the subgenus Pennaseius, in which the presence of only a single pair of preanal setae on the ventri-anal shield of the female, as well as the extreme length of setae s4, as shown by this species, are unique till now. up However, P. onilahy is also very similar to P. betsiboka, from which it differs, apart from the presence of setae Rl, in the absence of the second largest pore of the and in the postscutum size of the spermatheca. Phytoseius betsiboka and P. were onilahy placed according to the generally accepted division of the genus Phytoseius in different subgenera on the basis of the absence and presence, respectively, of the sublateral setae Rl. A natural relationship is not obvious as the two species do not fit the existing subgeneric division. Phytoseius betsiboka is in the nominate placed subgenus, but shows notocephalic pores on the dorsum which is typical for a majority of species of the subgenus Pennaseius. Description. Female: Dorsal shield smooth, 277 long and 160 Phytoseius onilahy is correctly classified with the latter subgenus, although with a species single wide; incised near seta r2; arrangement of pores similar to P. betsiboka, for absence of except second largest postscutal pore (fig. 9); fifteen pairs of setae on shield, the longer setae thickened, with rows of spines along their length; length: jl 30, j3 61, Z1 102, Z4 116, Z5 134, s3 28, s4 173, r2 54. Remaining dorsal setae (j4, j5, j6, J5, z4, z5, s2) smooth and minute, less than 6 long. Rl on interscutal membrane, 23 long. Peritremes reach nearly in front of setae j 1. pair of setae on the pre-anal ventri-anal shield are known only from the nominate subgenus. In addition to this intermingling of "subgeneric" characters, both species resemble each other so closely so as to form a natural species group, distinctly separate from all other known species of Phytoseius. In accordance with the still very imperfect knowledge on phytoseiid taxonomy, I have refrained from creating another subgenus, and have classified P. betsiboka and P. Sternal shield not well visible. Genital shield as usual, 75 wide. Ventri-anal shield 87 long and 55 onilahy conservatively, without regard of relationships.

6 One Five In very closely related to P. hongkongensis, P. minutus One Six There The - BIJDRAGENTOTDE DIERKUNDE, 46 (1) Etymology. P. onilahy derives its name from the river flowing South of Tulear. and P. kapuri. The setal dimensions of the Malagasy specimens are in almost complete agreement with those given by Van der Merwe (1968) for South African P. amba. Phytoseius (Pennaseius) hongkongensis Swirski & Shechter, 1961 Female: Setal dimensions: jl 19 25, j3 5056, Phytoseius (Phytoseius) hongkongensis Swirski & Shechter, Zl 7178, Z4 5965, Z5 6477, s3 1931, s , r2 3744, VL1 54; macrosetae genu 1961: 99, figs. 1 5; Denmark, 1966:44, fig. 17. Phytoseius (Pennaseius) hongkongensis; Ehara & Lee, 1971: 70, figs. 3237; Ehara, 1972: 169, fig. 81. IV 2327, tibia IV 3036, basitarsus IV 3034, distitarsus IV (A 9.25) collected on Pueraria javanica (Papilionaceae), Ivoloina, Tamatave, 7.VII One 9 (A 16.2) from the same plant in the same locality, 16.VII Iphiseius (Iphiseius) degenerans (Berlese, 1889) One <j (A 22.5) from Stylosanthes gracilis (Papilionaceae), 25.VII.1972 and seven 99 (A 27 series) from Solanum auriculatum also (Solanaceae),28.V1I.1972, near Ivoloina. Diagnosis. Denmark (1966) combined P. hongkongensis, P. amba Pritchard & Baker, 1962 and Seius degenerans Berlese, 1889 Iphiseius degenerans; Evans, 1954: 517, figs. 1 10; Athias- Henriot, 1957: 335, fig. 3B. Iphiseius (Iphiseius) degenerans; Pritchard & Baker, 1962: 299, figs. 6566; Van der Merwe, 1968: 105, figs P. minutus Narayanan, Kaur & Ghai, 1960, in the 9 (71.1), , one 9 (T 7.1), hongkongensis group of species. To this group, P. kapuri Gupta, 1969, should be added. In fact, these four forms are so similar, as to make their specific rank rather doubtful in my opinion. The lengths of some setae are the only distinguishing characters, and in this respect the specimens from the East Coast agree well with P. hongkongensis. 16.IX.1971, and two 99 (21.1 and 21.2), 9.XI.1971 from Fraxinus berlandieriana (Oleaceae) in Tananarive-Tsimbazaza. In the same locality, one (T 3.3) from Hibiscus rosa-sinensis (Malvaceae). 29.VII. 1971, one 9 (T 8.5) from an unidentified plant, 16.IX.1971, and one 9 (T 21.1) from lemon (Citrus limon; Rutaceae), 26.IX One <y (T 11.1) from papaya (Carica papaya; Caricaceae) in Tananarive-Nanisana, 21.IX.1971, and one 9 (T 17.1) from coffee (Coffea arabica; Rubiaceae) at Ambatomena, 30 km N.E. of Tananarive, 2.XII Female: Setal dimensions: jl 2327, j3 6268, zl 7782, Z4 7180, Z5 7073, s3 4245, s4 8796, r2 3743, VL1 4854; macrosetae genu Diagnosis. specimens are in perfect agreement with the available descriptions. IV 2227, tibia IV 2830, basitarsus IV 2225, distitarsus IV Iphiseius (Trochoseius) gongylus Pritchard & Baker, 1962 (figs. 1013) Remark. Blommers & Gutierrez (1975) this species was listed erroneously as P. amba. Iphiseius (Trochoseius) gongylus Pritchard & Baker, 1962: 304, figs Phytoseius (Pennaseius) amba Pritchard & Baker, and l ur <3 6 (series A 10) were collected from lemon leaves (Citrus limon; Rutaceae) at Ivoloina, Tamatave, 8.VII. 1972; one 9 (A 28.1) in the same locality, 29.VII.1972; three 99 and three <JcJ (A 29 series) Phytoseius (Pennaseius) amba Pritchard & Baker, 1962: 224, figs from combava (Citrus histrix) in the other specimens, 29.VII.I972. same plantation as the Phytoseius (Phytoseius) amba; Denmark, 1966: 46, fig. 18. Typhlodromus (Phytoseius) amba; Van der Merwe, 1968: 101, figs Diagnosis. can be little doubt about the identificationof this peculiar phytoseiid mite. 99 and one $ (17 series) from a not identified ornamental species of Malvaceae at Tananarive- Female: Large, bright red in life, in preparation Tsimbazaza, altitude 1250 m, 8.II One 9 (T 2.2) on Euphorbiapulcherrima (Euphorbiaceae), in the brownish. Dorsal shield 380 same locality, long and 335 wide. I8.VII Length of dorsal setae: jl 29, j3 54, Z1 18, Z4 (?) 225, s4 196, S2 214; remaining setae minute, less As Diagnosis. mentioned above, P. amba is than 10 long. Sublaterals r2 and R1 also small, about 8.

7 PHYTOSEIIDAE. 86 L. BLOMMERS - FROM MADAGASCAR Figs Iphiseius gongylus Pritchard & Baker, 1962: 10, spermatheca ; 11, peritremal plate ; 12, chelicera ; 13, chelicera. Figs Paraphytoseius multidentatus Swirski & Shechter, 1961: 14, chelicera ; 15. genital and ventri-anal shield. Figs Ambtyseius hima Pritchard & Baker, 1962: 16, genital and ventri-anal shield ; 17, spermatheca ; 18, chelicera ; 19, chelicera ; 20, ventri-anal shield

8 East BIJDRAGEN TOT DE - DIERKUNDE, 46 (1) Venter as figured with original description. (Pritchard & Baker, 1962) (Zaire), P. san- Ventri-anal shield 170 long and 230 wide. turcensis De Leon, 1965 (Puerto Rico), P. cras- Length tarsus IV 170. Length of macrosetae: centis (Corpuz & Rimando, 1966) (Philippines), P. on genu IV 43 and 102, tibia IV 70, basitarsus IV narayanani (Ehara, 1967) (India) and P. urumanus 62, genu III 60, tibia III 48, genu I 54. Chelicera in as fig. 12; fixed digit 20, movable digit 25 long. Spermatheca as in fig. 10; major duct 18, atrium 7, cervix 13 long. (Ehara, 1967) (Okinawa). The differences between the above are often slight, being mainly based on setal dimensions, and the number of specimens studied is low so that the specific status of several is questionable. Male: Dorsal shield 315 long and 260 wide. Length of long setae: jl and j3 25, Z4 205, s4 and S Genitosternal and ventri-anal shield reticulate as in female. Ventri-anal shield 170 long and 205 wide, not fused with peritremal shields, with three pairs of pre-anal setae, and one pair of pronounced pores at the same level as the posterior pair of pre-anal setae. Length tarsus IV 140. Length macrosetae on legs: genu IV 107, tibia IV 80, basitarsus IV 58, genu III 50, tibia III 45, genu I 50. Fixed digit of chelicera 16 long, movable digit 20. Major portion of spermatophoral process 12, branch 23. Paraphytoseius multidentatus Swirski & Shechter, 1961 (figs. 1415) As far as pertinent descriptions exist, my specimens agree best with P. multidentatus; the dimensions of the larger dorsal setae and the macrosetae of leg IV fall within the same range, except for dorsal seta Z5. Compared with P. urumanus, six setae are different, with P. santurcensis nine, and with P. eleven out of horrifer twelve. There exist some small differences between the specimens from the East coast and those from the West coast of Madagascar. The latter agree with the original description of P. multidentatus in having some small rod-like setae on leg IV and the macroseta on basitarsus IV considerably longer than on distitarsus IV. The specimens from Tamatave, however, have all setae, except the macrosetae and one small seta on the femur, acuminate and both macrosetae on tarsus IV of nearly the same length, as in the topotype series Paraphytoseius figs. 7, multidentatus Swirski & Shechter, 1961: 114, described by Ehara & Lee (1971). Amblyseius (Paraphytoseius) multidentatus; Ehara & Lee, 1971 Female: Dorsal shield long and , figs coast: 39 9 (T 14 series) were collected 160 wide; with at least twenty pairs of pores, those near setae z5 very pronounced. Dimensions from Phaseolus lunatus (Papilionaceae) and I 9 (T 15.1) from of the larger, serrate setae: jl 3036, j3 7989, Sechium edule (Cucurbitaceae) at Bains des Dames, Tamatave City, 18.X In the same locality, 1 9 (A 11.1) from P. lunatus, 12.VII At Ivoloina, Tamatave, 3 99 and ' 3 (A 6 series) were found on P. lunatus, 11.II In the same locality on Pueraria javanica (Papilionaceae), and 3 <J (A 9 series), 7.VII.1972, 299 and 1 <J (A 16 series), 16.VII.1972, and and 2 <5 (J (A 30 series), 1.VIII West coast: II 9 9 (M 10 series) were collected from Urena lobata (Malvaceae), I 9 (M 18.4) from Alchornea sp. (Euphorbiaceae) and 1 <y (M 20.3) from Phaseolus sp., all at Ambato Boeni, 24.VI On the same day, (M 15 series) were found on Leonotis nepetaefolia (Labiatae) at Ampijoroa, Tsaramandroso. On 9 and 24.III.1971, a"d 4 (J (J (47 series) were collected from Sida sp. (Malvaceae) in the botanical garden of the Agricultural Station at Tulear. On 25.IV. 1971, and 2 (J (J (B 13 series) were found on the same plants in the same place. Z4 6880, Z , s , r2 3648, R Remaining setae smooth and less than 10 long. Peritremes reach in front of setae jl- Venter as described by Ehara & Lee (1971). Ventri-anal shield long and 5963 wide, with a pair of minute pores. VL long. Leg IV with four distinct macrosetae: on genu 2227, tibia 2838, basitarsus 3845 and distitarsus In the specimens from the East coast, all remaining setae on the legs acuminate, except the most proximal seta on femur IV, and one seta on genu II, which are both rod-shaped. Diagnosis. Only six species of the genus Paraphytoseius Swirski & Shechter, 1961, have been described: P. multidentatus (Hong Kong), P. rifer hor- The specimens of the West coast with a rodshaped seta on femur IV, genu IV (often two), tibia IV, basitarsus IV and genu II. Spermatheca as usual in Paraphytoseius ; long

9 - 88 L. BLOMMERS PHYTOSEIIDAE FROM MADAGASCAR female. Length of macrosetae: genu IV 20, tibia IV 30, basitarsus IV 33, distitarsus IV Chelicera as figured (after A 9.24). The pointed structure halfway the spermatophoral process seems less pronounced in the West coast Fixed digit of chelicera with about eight teeth, and slender major duct (20 x 2), knob-like atrium, platter-shaped cervix, about 7 in diameter; vesicle filled with spermatophore, globular, about 30 in diameter. Spermatophore itself pearshaped. specimens. Length movable digit with three. of both digits about 20, spermatophoral process 17. Fixed digit with about eight Male: Peritremes shorter than in the female, not teeth, movable digit with one. reaching as far as base of setae j3. Dorsal shield 220 long and 140 wide, incised near setae s4 as in the female. Large pores near setae z5. Length of Field notes. This species was observed frequently on plants the larger serrate setae: j 1 27, j3 55, Z4 48, Z5 60, r2 30, R1 13. The latter two located on the shield. which harboured also the tetranychid mite species Tetranychus macfarlanei Baker & Pritchard, Remaining dorsal setae smooth and minute (T 14, T 15, A 6, A 11, A 30) and Three pairs of pre-anal setae and one pair of pores on ventri-anal shield. VL1 22 long. Macrosetae and rod-like setae on legs as in Eotetranychus limoni Blommers & Gutierrez, 1975 (A 9) in the region of Tamatave, and T. neocaledonicus Andre, 1933 (B 13) in Tulear. Unsuccess- Fig. 21 Amblyseiushima Pritchard & Baker, 1962: dorsum. Figs Amblyseius hova sp. n.: 22, chelicera ; 23, venter.

10 At The Holotype A. BIJDRAGEN TOT DE - DIERKUNDE, 46 (1) ful attempts to rear P. multidentatus using the Field notes. latter spider mites as prey, suggest that this The plants on which this phytoseiid was found phytoseiid mites. mite is not a predator of Tetranychus often harboured considerable numbers of the tenuipalpid Brevipalpus sp. (T 1, T 5, T 6) as well Amblyseius (Amblyseius) hima Pritchard & Baker, 1962(figs ) as tydeid mites and scale insects (T 1, T 5). Amblyseius (Amblyseius) hova sp. n. (figs. 2226) Amblyseius (Amblyseius) hima Pritchard & Baker, 1962: 257, figs and 1 (j Tsimbazaza, Tananarive (alt m), (series T 1) were collected on leaves of Euphorbia pulcherrima (Euphorbiaceae), 18.VII.1971; and 3 $ 3 (series T 5) on Thunbergia grandiflora (Acanthaceae), 7.IX.1971; 1 9 (T 6.3) on Barleria sp. (Acanthaceae), 16.IX.1971; 3 99 and 1 on <y (series 20) Dombeya sp. (Sterculiaceae) in January (T 3.5) and 1 paratype 9 (T 3.4) from Hibiscus rosa-sinensis Tananarive- (Malvaceae), Tsimbazaza, alt m, 29.VII.1971; 3 paratypes (9 9) (T 6 series) from Barleria sp. (Acanthaceae) and 1 paratype (9) (T 8.10) from an unidentified plant, both in the same locality as the holotype, 16.IX Two young, not yet fully sclerotized 9 9 (T 10A series) collected on lemon (Citrus limon; Rutaceae), 1 9 (T 11.2) on papaya (Carica papaya; Caricaceae), both in Tananarive-Nanisana, 21.IX.1971, and 1 9 Diagnosis. original description is based on specimens from Ruanda Urundi, and the Malagasy specimens, though of smaller size, agree well with the originals. on (T 5.13) Thunbergia grandiflora (Acanthaceae), Tsimbazaza, 7.IX.1971 belong to the same species. Differential diagnosis. hova bears a close resemblance to South African A. munsteriensis Female; Dorsal shield imbricated, long, Van der Merwe, It differs markedly from wide; with at least fifteen pairs of this species in the shape of the spermatheca. pores. Length dorsal setae: j , j3 27, j4 20, j5 1820, j6 1820, J2 2021, J5 56, z4 3032, z5 1820, Z1 2021, Z4 2325, Z5 4550, s2 2729, s4 3436, S2 2527, S4 2325, S5 2527, r2 21, R1 18. Peritremes do not reach much further than sublaterals r2. Sternal shield posteriorly with median lobe. Ventri-anal shield agrees with original description; 8286 long and 4654 wide. Surrounding membrane with six pairs of pores and four pairs of setae; VL long. One pair of metapodal platelets. Tarsus IV 100 long, macrosetae on it Spermatheca as figured, major duct 11, cervix 20 long; atrium 3, vesicle 19 in diameter. Digits of chelicera 22 long. Male: Dorsal shield anteriorly imbricated, 230 long and 135 wide. Length of setae: jl 21, j3 25, j4 18, j5 16, j6 18, J2 16, J5 5, z4 26, z5 18, Z1 18, Z4 20, Z5 40, s2 24, s4 29, S2 20, S4 20, S5 22, r2 22, R1 16. Sublaterals r2 and R1 on dorsal shield. Peritremes as in female. Ventri-anal shield 86 long, not fused with peritremal shield; only two pairs of pores visible; three pairs of pre-anals. VL1 23. Macroseta on basitarsus IV 32 long. Description. Female: Dorsal shield rather strongly imbricated; 340 long and 215 wide, with at least thirteen pairs of pores. Seventeen pairs of setae; length: jl 23, j3 25, j4 12, j5 11, j6 13, J2 14, J5 9, z4 20, z5 13, Z1 14, Z4 20, Z5 72, s2 20, s4 23, S2 20, S4 22, S5 22. Z5 smooth. Sublaterals r2 and RKon interscutal membrane, 16 and 12 long, respectively. Peritremes reach in front of setae j 1. Posterior margin of sternal shield W-shaped. Genital shield as usual. Ventri-anal shield 104 long and 77 wide, with three pairs of pre-anals. Surrounding membrane with six pairs of pores, and four pairs of setae; VL1 42 long. Length tarsus IV 116. Macrosetae on IV genu 36, tibia IV 30, basitarsus IV 86, genu III 25 long. No other macrosetae present. Chaetotaxy of legs normal. Chelicera not easily discernable in type series, about 25 long. Fig. 22 is drawn after specimen T 10A5. Spermatheca small, cervix funnel-shaped, 12 long. Male: Unknown Field notes. Various species Chelicera as figured, digits 18 long, spermatophoral process gibbet-shaped. Major portion 18 long, branch 9. of small arthropods were observed in the direct vicinity of A. hova, such as Brevipalpus sp. (T 5, T 6, T 10) and Tetranychus

11 PHYTOSEIIDAE One These 90 L. BLOMMERS - FROM MADAGASCAR neocaledonicus (T 3, T 10), unidentified Ty- 9 (M 32.1) collected on Hibiscus sp. deidae and white flies (Aleurodidae) (both T 3), but there was no indication that the phytoseiid uses any of them for food. (Malvaceae) at Ampijoroa (Majunga province), 30.XI Four 9 9 and four c? (J (M 22-2k series) from a mass rearing started with specimens found on several herbs near Katsepy (Bombetoka Bay, opposite Majunga), 27.IV Etymology. The Hova were of old the free men of the Merina, inhabitants of the highlands around Diagnosis. specimens the type series from Tulear. are identical to Tananarive. Amblyseius (Amblyseius) bibens Blommers, 1973 Amblyseius (Amblyseius) masiaka Blommers & Chazeau, 1974 Amblyseius (Amblyseius) bibens Blommers, 1973: 111 figs Amblyseius (Amblyseius) masiaka Blommers & Chazeau, 308, figs : Tulear: (B 3 series) from Carica papaya (Caricaceae), 3.III.1971; 8 99 and2<j(s (B 4 and 40 series) Figs Amblyseius hova sp. n.: 24, dorsum ; 25, spermatheca ; 26, leg IV. Fig. 27. Amblyseius ankaratrae sp. n.: leg IV.

12 BIJDRAGEN TOT DE DIERKUNDE, (1) 91 from Sida spinosa (Malvaceae), ; (B 6 series) from Sida sp., 6.III.1971; 299 (B 7 series) from Leonotis and 1 (5 (M 11 series) from Plumeria alba (Apocynaceae) at Ambato Boeni on the same day. nepetaefolia (Labiatae), 4.III.1971; (B 15 series) from Corchorus trilocularis (Tiliaceae), ; 699(8 23 and 57 series), 69 9 (62 series) and 299 and 1 <J (65 series) Amblyseius (Amblyseius) brevipes Blommers, 1973 from the same plant species on , l and 2.1 V.1971, respectively; 6 99 (B 22 and 61 series) from Corchorus Amblyseius (Amblyseius) brevipes Blommers, 1973: 112, figs. olitorius, ; (B 24 series) from Abelmoschus esculentus (Malvaceae), 25.III.1971; (B 34 series) from cotton (Gossypium hirsutum; Malvaceae), ; 299 Tulear: except for in the type locality, this from Manihot utilissima (Euphorbiaceae), 15.V Tananarive: and 1 5 (T 25 series) from Phaseolus vulgaris (Papilionaceae) at Tananarive-Soanierana (alt m)j4.iii.i973. Tamatave: Apart from the specimens recorded previously from this region (Blommers, 1974a), this species was also encountered at Bains des Dames, Tamatave City, on Phaseolus lunatus (Papilionaceae) ( 3 99 ar>d 1 <3; T 14 series), Sechium edule (Cucurbitaceae) (3 9 9; T 15 series) and species (1 9; 52.8) was also collected on Acalypha sp. (Euphorbiaceae), 8.III Ambato Boeni/Ampijoroa: (M 17 series) from Carica papaya (Caricaceae) in Ampijoroa, 24.IV.1972; (M 20 series) from Phaseolus (?) sp. near Ambato Boeni on the same date as the first; 1 9 (M 26.21) from Alchornea sp. (Euphorbiaceae) in Ambato Boeni, 12.IX Tamatave: an <3 1 <J (A 4 series) from C. papaya at the l.f.a.c.-station in Ivoloina, 8.II Ricinus communis (Euphorbiaceae) (2 9 9; 18.X T '6 series), Tananarive; (T 6 series) from Barleria sp. (Acanthaceae) at Tsimbazaza (1250 m), 16.IX.1971; on the same place at the same date (T 7 series) from Fraxinus berlandieriana (Oleaceae) and and 1 cj (T 8 series) from the Field notes. This predacious mite was encountered always in undergrowth under these trees. At Tananarive-Nanisana, 1 9 (T 11.4) was collected on C. papaya, 21.IX or near populations of spider mites of the genus Tetranychus: T. neocaledonicus in the region of Tulear, T. macfarlanei near Tamatave and T. urticae Koch, 1836, and T. ludeni Zacher, 1913, in the central highlands. Discussion. Amblyseius rotundus and A. brevipes have been found to be more intimately related than realized at the time of their original description. The original diagnosis appeared to be insufficient to Amblyseius (Amblyseius) rotundus discriminate between them over their whole area Blommers, 1973 of occurrence. The following biometrical study is Amblyseius (Amblyseius) rotundus Blommers, 1973: , figs. meant to provide these means. Other morphological characters such as the shape of the spermatheca and of the chelicera were not taken into Tulear: an<i ' <5 (B 9 and 66 series) from consideration as they are difficult to quantify. We Mangifera indica (Anacardiaceae), ; 599(8 10 and 52 series) from Acalypha sp. (Euphorbiaceae), 8.III.1971; (B 20 and 55 series) from Croton sp. (Euphorbiaceae), ; 4 99 (56 series) from Echinochloa colonna have measured the longer setae, notably those on leg IV, and the length of the same leg (genu, tibia, basi- and distitarsus), since these constituted also (Gramineae), ;699 and 3 <}<} (57 series) from the major distinguishing marks between the two Corchorus trilocularis (Tiliaceae), ; 299(8 24 series) from Abelmoschus esculentus (Malvaceae), ; from the same plant and 1 S (60 series), 6.1 V.1971; I<} (B 29.1) from Sida sp. (Malvaceae), 6.1V.1971; 19 (B 33.4) from Acalypha sp. (Euphorbiaceae), 13.IV. 1971; (B 34 series) from cotton (Gossypium hirsutum; Malvaceae), 20.1V. 1971; 299 and 1 $ (B 35 series) from Bauhinia sp. (Caesalpinaceae), 11.IV.1971; and 69 9 fr m manioc (Manihot utilissima; Euphorbiaceae), 15.V Majunga-Katsepy: 1 9 (M 1.3) from unidentified Asclepiadaceae, 26.IV.1971; 1 9 ar d 1 5 (M 3 series) from an unknown species according to the original description. In table I and fig. 65 (p. 102) the measurements are combined in seven groups. The data show a clear difference between A. rotundus and A. in brevipes the region of Tulear, their type locality. Evidently, the specimens from Tamatave (all from C. papaya) and two samples from Ambato Boeni/Ampijoroa (from C. papaya and Phaseolus sp.) belong also to A. brevipes. On the plant; I other hand, the animals collected on the dune 9 (M 7.1) from Rhopalocarpus lucidus (Rhopalocarpaceae); 499 and 1 <} (M 9 series) from Bauduinia sp. (Caesalpinaceae) and 799 and 2 $$ (M 16 series) from Diospyros sp. (Ebenaceae). All collected on the same date as the first. Ambato Boeni/Ampijoroa: 3 9$ (M 6 series) from Ricinus communis (Euphorbiaceae) at Ampijoroa, 24.IV. 1975; vegetation in Majunga/Katsepy are certainly A. rotundus. The two remaining groups are more difficult to assign. The other two samples (from Plumeriaand from Ricinus) from the region of Ambato Boeni/

13 Holotype The 92 L. - BLOMMERS PHYTOSEIIDAE FROM MADAGASCAR Table I. Size and standard - (mean deviation) of leg IV (femur distitarsus) and the macrosetae on genu and basitarsus of the same leg in grouped female specimens in the Amblyseius rotundus-brevipes complex. seta genu IV seta basitarsus IV length leg IV length leg IV N No. samples seta genu IV (= plants) x SD x SD x SD A.Tulear brevipes B. Ambato Boeni/Ampijoroabrevipes C. Tamatave , D.Tananarive E. Ambato Boeni/Ampijoroarotundus F. Tulear rotundus G. Majunga Katsepy Ampijoroa are sufficiently different from those Sternal and genital shield as usual. Ventri-anal identified as A. brevipes from the same region, to shield 100 long, 75 wide; widest on the level of be named A. rotundus. Certainly so, if one knows paranal setae; with three pairs of pre-anal setae. that in Ampijoroa Carica papaya (with brevipes) and Ricinus communis (with rotundus) grew side by side. The specimens from Tananarive, finally, have rather long macrosetae, but also the longest legs of all specimens examined. If this latter dimension is considered as a measure for the body size, Surrounding membrane with four pairs of pores and four pairs of setae; VL1 18 long. Length tarsus IV 128. Length of macrosetae: on genu IV 125, tibia IV 84, basitarsus IV 32, genu III 38, tibia III 32, genu II 30, genu I 36. Fixed digit of chelicera 34 long, with three blunt subapical teeth and eight in row. Movable which itself is difficult to determine in mounted digit 34, without teeth. specimens, the relative lengths ofthe macrosetae are entirely comparable with those found with the other A. brevipes (see table I). Therefore, we are strongly inclined to assign them to this species. Spermatheca as in fig. 29. Major duct 16, cervix 7 long. Male: Dorsal shield285 long, 180 wide. Length of setae: jl 20, j3 29, j4 5, j5 5, j6 5, J2 9, J5 7, z4 7, z5 5, Z1 11, Z4 9, Z5 170, s2 9, s4 13, S2 9, S4 9, S5 7, R1 9. Amblyseius (Amblyseius) ankaratrae sp. n. (figs ) Ventri-anal shield imbricated, 115 long; probably fused with peritremal shields; with four preanals and at least four pairs of pores. 9 (T 4.1) and four paratypes (3 9$ and I <J) from Dicoryphe sp. (Hamamelidaceae) in mountain rain forest, Manjakatompo Forest Station, Ankaratra Mountains, alt m, 14.VIII Length of macrosetae: on genu IV 70, on tibia IV 64. Chelicera in our specimen not entirely visible (fig. 32); length digits 22; fixed digit with one Differential diagnosis. combination of an elongate, whip-like seta Z5 with a short Z4 is unique the known among species of Amblyseius. subapical tooth and seven in row. Spermatophoral process gibbet-shaped; major portion 18, branch 7. Description. Female: Dorsal shield smooth, with at least twelve pairs of pores; 350 long and 250 wide. Seventeen pairs of setae; length: jl 20, j3 27, j4 5, j5 7, j6 9, J2 11, J5 5, z4 U,z5 7, Z1 13, Z4 14, Z5 205, s2 11, s4 13, S2 11, S4 11, S5 11. Sublaterals r2 and R1 on interscutal membrane, 13 and 11 Amblyseius (Amblyseius) boina sp. n. (figs ) and I Holotype 9 (M 18.6) and seven paratypes (699 (J) were collected on Alchornea sp. (Euphorbiaceae) in degraded savannah brush near Ambato Boeni (Majunga province), 24.1 V long. Peritremes reaching base of setae jl. A Differential diagnosis. This species differs finely striated band the bordering dorsal scutum from most other Amblyseius species is clearly visible in mounted specimens. in the combined presence of long whip-like setae Z4 and

14 BIJDRAGEN TOT - DE DIERKUNDE, 46 (1) Figs Amblyseius ankaratrae sp. n.: 28, dorsum ; 29, spermatheca ; 30, chelicera ; 31, venter ; 32, chelicera.

15 Nine Two These 94 L. BLOMMERS - PHYTOSEIIDAE FROM MADAGASCAR Z5, and a feebly pronounced corniform atrium. In this respect, A. boina is closely related to A. Amblyseius (Amblyseius) tamatavensis Blommers, 1974 aerialis (Muma, 1955), A. paraaerialis Muma, 1967, A. lassus Schuster, 1966, and A. invictus Amblyseius (Amblyseius) tamatavensis Biommers, 1974a: 144. Schuster, However, A. aerialis and A. lassus figs have a more tubular cervix of the spermatheca, A. paraaerialis has considerably shorter setae Z4, Z5 and s4, and A. invictus has a knob-like atrium of the spermatheca and a more triangular ventrianal shield. 9 9 a"d nine $ c} (M 22-2k series) from a mass rearing starting with specimens collected on several plants (Phaseolus sp., Papilionaceae; Leonotis sp., Labiatae, and Ipomoea sp., Convolvulaceae) near Katsepy, Bombetoka Bay opposite Majunga, 27.IV Description. Female: Dorsal shield smooth, 320 long and 255 wide; with at least fourteen pairs of pores. Diagnosis. type series from Tamatave. specimens are similar to the Seventeen pairs of setae; length: jl 29, j3 41, j4 5, j5 4, j6 5, J2 5, J5 7, z4 7, z5 5, Z1 7, Z4 114, Z5 236, s2 7, s4 80, S2 7, S4 8, S5 8. Sublaterals r2 and R1 13 and 9 long, respectively. Peritremes reach in front of setae jl. Sternal and genital shield as usual. Ventri-anal shield rather narrow, 103 long and 75 wide, with three pairs of pre-anal setae. Surrounding membrane with five pairs of pores and four pairs of setae; VL1 75 long. Tarsus IV 120 long. Length of macrosetae on legs: genu IV 110, tibia IV 84, basitarsus IV 82, genu III 59, tibia III 42, genu II 40, genu I 45. Fixed digit of chelicera 25 long, with two subapical teeth, and nine in a row. Movable digit Amblyseius (Amblyseius) deleoni Muma& Denmark, 1970 Amblyseius (Amblyseius) largoensis; Muma, 1961 (nec largoensis Muma, 1955); Schuster & Pritchard, 1963: 237, fig. 26; Muma, 1964: 22; Van der Merwe, 1968: 135, figs Amblyseius deleoni Muma & Denmark, 1970: 68, figs (T 3 series) collected from Hibiscus rosa-sinensis (Malvaceae), 29.VII.I971; ten 9 9 (T 6 series) from Barleria sp. (Acanthaceae) and three 9 9 (T 7 series) from Fraxinus berlandieriana (Oleaceae), both on 16.IX.1971; twelve 9 9 (T 9 series) from Ageratum conyzoides (Compositae) on 17.IX All these specimens were collected in the zoological garden of the O.R.S.T.O.M. centre in Tananarive- Tsimbazaza, at 1250 m altitude. 25 long, with three teeth. Spermatheca as figured, feebly sclerotized, 45 long. Major duct 22 long and 1 wide, atrium illdefined, cervix 13 long. Male: Dorsal shield 275 long, 178 wide. Length of setae: jl 21, j3 36, j4 4, j5 4, j6 5, J2 5, J5 5, z4 9, z5 4, Z1 5, Z4 80, Z5 180, s2 7, s4 63, S2 7, S4 5, S5 5, r2 9, R1 5. Ventri-anal shield 98 long, fused with peritremal shields, with four pairs of pores and four pairs of pre-anals. VL1 45 long. Length of macrosetae on legs: on genu IV 77, Diagnosis. According to Muma & Denmark (1970) the name A. largoensis had been used for two different since Muma species (1961), and A. largoensis Muma, 1961, was consequently renamed A. deleoni. I have compared the Malagasy specimens with a specimen of A. deleoni from Florida identified and sent to me by Dr. H. Denmark. They are very probably conspecific, while are certain- they ly identical to A. largoensis Muma, Merwe from (1968) South Africa. sensu Van der tibia IV 54, basitarsus IV 63, genu III 36, 30, genu II 29, genu I 32. tibia III Setal dimensions in the female: jl 34-38, j , Z , Z , s ; length of Length of both digits of chelicera 18. Fixed macrosetae: on genu IV , tibia IV 80-86, digit with one subapical tooth and eight (or seven) in a row. Movable digit with one tooth; basitarsus IV 68-75; s4 is nearly always shorter than Z4. Z4 often faintly serrate. slightly spermatophoral process portion 13 long, branch 5. as in fig. 38, major We failed to observe males in the mass rearing (see below) of this species, and eggs reared singly produced reproducing females. This thelytoky Etymology. was also observed by Van der Merwe (1968), but Boina is called the western lowland region on males are mentioned by Muma & Denmark both sides of the Betsiboka River. (1970).

16 BIJDRAOEN TOT DE DIERKUNDE, 46 - (1) Figs Amblyseius boina sp. n.: 33, dorsum ; 34, chelicera ; 35, leg IV ; 36, spermatheca ; 37, ventri-anal shield ; 38, chelicera ; 39, genital and ventri-anal shield.

17 96 L. - BLOMMERS PHYTOSEIIDAE FROM MADAGASCAR Field notes. Length tarsus IV (including basitarsus) 160. A. deleoni was found in company of all sorts of Five macrosetae on leg IV: two on genu 128 and small arthropods, notably the tenuipalpid mite 36, two on tibia 104 and 27, one on basitarsus 70. Brevipalpus sp. (T 6, T 7, T 9), white flies (T 3, T 9), tydeid mites (T 3, T 7) and low numbers of Tetranychus neocaledonicus (T 3, T 9). Length of remaining macrosetae: genu III 50, tibia III 45, tarsus III 36, genu II 38, genu I 41. Genu II with seven pairs of setae. Fixed digit of chelicera with two subapical teeth and eight in a row. Movable digit with three tiny teeth. Length of both digits about33. Spermatheca as figured. Length cervix 36. Male: Dorsal shield 310 long and 220 wide; r2 and R1 on dorsal shield. Length of setae: jl 34, j3 48, j4 5, j5 4, j6 7, J2 7, J5 9, z4 9, z5 5, Z1 9, Z4 80, Z5 218, s2 12, s4 86, S2 13, S4 11, S5 9, r2 14, R1 11. Ventri-anal shield imbricate, with three pairs of pores, fused with peritremal shields, 128 long. Macrosetae on IV 86 legs: genu and 29, tibiaiv 73 and 21, basitarsus IV 63, III genu 38, tibia III 36, basitarsus III 32, genu II 34, tibia II 36, genu I 32. Length tarsus IV 130. Fixed of chelicera with digit one subapical tooth and six teeth in a row. Movable digit with one tooth; spermatophoral as process figured, major portion 10 long, branch 5. Both digits 20 long. Amblyseius (Amblyseius) sakalava sp. n. (figs ) Holotype 9 (65.3) and 5 ar paratypes (3 >d (J <J; series 65) collected on Corchorus trilocularis (Tiliaceae) in the Agricultural Garden in Tulear, 2.1V Other paratypes: and 2 S6 (52 series) from Acalypha sp. (Euphorbiaceae) and I 9 (49.1) from Lagerstroemia indica (Lythraceae) in the same locality as the holotype, 8.III Specimens (3 9 9 ar) d 2 g<$, B 24 series) were also found on okra, Abelmoschus esculentus (Malvaceae) in the same garden, 25.III.1971, and on cassava (Manihot utilissima; Euphorbiaceae), 15.V Differential diagnosis. A. sakalava compares well with both A. largoensis (Muma, 1955) and A. neolargoensis Van der Merwe, It differs from the latter species in the considerably smaller size of the dorsal setae s4, Z4 and Z5. A. largoensis was never described in sufficient detail, but through the courtesy of Dr. H. Denmark I am able to compare my specimens with a specimen of A. largoensis from Florida; in A. sakalava all whip-like setae (also on the legs), except dorsal Z5, are definitely longer than in the American A. largoensis, which is also different in having only 2 chelicera. + 6 teeth on the fixed digit of the The setal lengths of A. sakalava well with agree those described by Ehara (1959) for A. largoensis Field note. Tetranychus neocaledonicus was always present and often quite numerous on the plants from which A. sakalava was recovered. Etymology. The Sakalava people inhabit the entire western coast and lowlands from Ambanja in the North to from Japan, but since the spermatheca was not Tulear in the South. described the latter might have been A. deleoni just as well. Amblyseius (Amblyseius) trichophilus sp. n. Description. Female: Dorsal shield smooth, nearly 400 long (figs ) and 270 wide; with at least fifteen pairs of pores; seventeen pairs of setae: j 1 43, j3 57, j4 7, j5 4, j6 9, J2 11, J5 11, z4 9, z5 5, Z1 11, Z4 111, Z5 264, s2 13, s4 104, S2 20, S4 14, S5 12. Sublaterals on interscutal membrane; r2 14 and R1 10 long. Peritremes reaching in frontof setae j 1. Sternal shield with straight posterior border and genital shield as usual. Ventri-anal shield elongate and constricted; length Holotype 9 (M 12.1) and two paratypes (M 12 series; 299) from Pueraria javanica (Papilionaceae),Ambato Boeni, 24.IV.1972; 3 paratypes (M 15 series; 299 and 1 <J) from Leonotis nepetaefolia (Labiatae), Tsaramandroso, Ampijoroa, on the same day as the holotype; 5 paratypes (M 28; and 3 <3cJ) from Combretum villosum (Combretaceae), Route Nationale 4, km 491, Amboromalandy, II.IX.1972; and 2 paratypes (M 34 series; 9 and (?) from Leptadenia madagascariensis (Asclepiadaceae), Tsaramandroso, Ampijoroa, 30.XI One 9 (M 27k7) was collected on Urena lobata (Malvaceae), Tsaramandroso, Andranofasika, 12.IX. 128 and maximal width 80. Four pairs of both pores and setae on surrounding membrane; VL1 70 long One 9 f rom cassava (Manihot utilissima; Euphorbiaceae), Tulear, 18.V. 1971,belongs to the same species.

18 BIJDRAGEN TOT DE DIERKUNDE, - 46 (1) Figs Amblyseius sakalava sp. n.: 40, dorsum ; 41, spermatheca ; 42, leg IV ; 43, chelicera ; 44, genital and ventri-anal shield ; 45, chelicera ; 46, ventri-anal shield.

19 98 L. - BLOMMERS PHYTOSETIDAE FROM MADAGASCAR Figs Amblyseius trichophilus sp. n.: 47, dorsum ; 48, leg IV ; 49, chelicera ; 50, spermatheca ; 51, genital and ventri-anal shield ; 52, ventri-anal shield ; 53, chelicera. Figs Amblyseius sundi Pritchard & Baker, 1962: 54, spermatheca ; 55, chelicera ; 56, chelicera, spermatophoral process viewed laterally.

20 A. Nine Some The BIJDRAGEN - TOT DE DIERKUNDE, 46 (1) Differential diagnosis. trichophilus is diffi- Remark. specimens are substantially cult to compare with known members of the genus. The dorsal setal pattern places it clearly within the nominate subgenus, but several other characters of this are rare or even species unique among members of this group. In fact, except for larger than the holotype, notably those of the M 28 series. Some measurements on 9 M 28.1 might illustrate this: dorsal shield 310 long and 200 wide, jl 23, j3 54, z4 59, Z4 70, Z5 70, s4 64, S2 70. the presence of 17 pairs of dorsal setae, A. trichophilus resembles in almost every important feature members of the subgenus Proprioseius Chant, The possession of a number of longer serrate dorsal setae, the absence of distinct macrosetae on leg IV, the elongate form of the ventri-anal shield in the female, and the Field notes. It should be stressed that A. trichophilus was found often on plants infested with spider mites of the genus Eotetranychus, viz., E. paracybelus Gutierrez, 1967 (M 27), E. savanae Gutierrez, 1967 (M 34) and an undescribed third species (M 12). shape of the spermatheca, are characters common in Proprioseius but rare in Amblyseius. Amblyseius (Proprioseiopsis) sundi Description. Pritchard & Baker, 1962, comb. n. (figs ) Female: Small-sized species. Dorsal shield 275 long and 150 wide, smooth, with at least nine pairs of pores, of which four large and conspicuous. Seventeen pairs of setae, length: jl 23, j3 41, j4 11, j5 11, j6 11, J2 11, J5 7, z4 39, z5 7,Z1 9, Z4 57, Z5 57, s2 21, s4 54, S2 54, S4 20, S5 11. Setae jl, j3, z4, Z4, Z5, s2, s4, S2 and r2 distinctly serrate. Sublaterals r2 32 and R1 20 long. Peritremes reaching level with setae jl. Sternal shield with straight posterior margin. Genital shield as usual. Ventri-anal shield slender, 90 long and 50 wide; three pairs of pre-anals. Pre-anal pores absent. Surrounding membrane with six pairs of pores and four pairs of setae; VL1 38 long. Legs without pronounced macrosetae. Genu II with 8 setae; other segments and legs normal. Both digits of chelicera 30 long. Fixed digit with two subapical teeth and tooth one halfway. Movable digit with one tiny tooth. Spermatheca as figured. Major duct 20 long. Atrium knob-like. Cervix platter-shaped. Amblyseius (Amblyseius) sundi Pritchard & Baker, 1962: 244, figs (Rutaceae) in 9 9 (T 10 series) collected on Citrus limon the Agricultural Station at Nanisana, Tananarive, alt m, 21.IX.1971, and three 9 9 and five <5 cj (T 10k series) from a mass rearing started with specimens of the same origin. Four 9 9 and one (M 6k series) originate from a rearing begun with specimens found on Ricinus communis (Euphorbiaceae) near Tsaramandroso, Ampijoroa, 24.IV.I972. Diagnosis. absence of setae Zl, which places this species clearly into the subgenus Proprioseiopsis Muma, 1961, and the extreme length of the dorsal setae Z4, Z5 and s4 leave little doubt about the identity of our specimens. Only minor differences in setal lengths and the existence of males distinguish them from thelytokous A. parasundi Blommers, 1974, reported previously from the East coast (Blommers, 1974a). Female: Dorsal shield long and wide. Length of setae: jl 37-45, j , Z Male: Dorsal shield 220 long and 125 wide. Length of setae jl 18, j3 25, j4 12, j5 14, j6 14, J2 214, Z , s , r All remaining setae minute, less than 10 long. 12, J5 5, z4 36, z5 9, Z1 12, Z4 37, Z5 34, s2 20, s4 40, S2 40, S4 14, S5 11, r2 25, R1 16. Longer setae serrate as in female. Ventri-anal shield fused with peritremal Venter agrees with original description in Ventri-anal shield shape long and wide. VL Length of macrosetae: on genu IV and shields, 100 long, with four pairs of pre-anals; 54-66, tibia IV and 46-54, basitarsus IV three pairs of pores; VL1 18 long and 28-29, genu III 70-89, tibia III 59-71, Legs without macrosetae. basitarsus III 37-48, genu II 45-53, genu I Digits chelicera 18 long. Fixed digit with two Two more whip-like macrosetae on both tibia teeth, movable with one. Spermatophoral process gibbet-shaped, major portion 12 long, branch 3. and tarsus I. Length of tarsus IV (including basitarsus)

21 Field note. One The 100 L. - BLOMMERS PHYTOSEIIDAE FROM MADAGASCAR Both digits of chelicera about 40 long. Fixed nineteen pairs of pores. Sixteen pairs of setae digit with two subapical teeth and 9-11 in a row. Movable digit with three teeth. Spermatheca with cervix long. Male: Dorsal shield x Length of setae: jl 30, j3 47, Z4 143, Z5 364, s Venter as usual. Ventri-anal shield fused with peritremal shields, 120 long. Macrosetae on legs: genu IV 160 and 40, tibia IV 125 and 40, basitarsus IV 98 and 20, genu III tibia 55, III 46, tarsus III 32, genu II 38 and genu I 55. Chelicera as figured. Length digits 22. Major portion of spermatophoral process 14, branch 9. Field notes. Some Tetranychus neocaledonicus and Brevipalpus sp. were observed on the lemon trees on which this species was collected. Ricinus communis in Ampijoroa contained rather large numbers of the same spider mite. (pair J2 lacking); length: j 1 20, j3 27, j4 3, j5 4, j6 4, J5 7, z4 9, z5 4, Z1 9, Z4 60, Z5 104, s2 13, s4 54, S2 9, S4 9, S5 9. Sublaterals r2 and R1 both 12 long. Peritremes reaching in front of setae jl. Sternal shield with straight posterior margin. Genital shield as usual. Ventri-anal shield 115 long and 90 wide, mildly imbricated, pairs of pre-anals. Surrounding with three membrane with three pairs of pores and four pairs of setae; VL1 62 long. Tarsus IV 114 long. Length of macrosetae: on genu IV 48, tibia IV 30, basitarsus IV 61, genu III 20, genu II 20. Genu II with eight setae (VHI-type according to Van der Merwe, 1968). Both digits of chelicera 28 long. Fixed digit with two subapical teeth and seven in row. Movable digit with a single tiny tooth. Major duct spermatheca 21 long, constricted in the middle; atrium small; cervix caliciform, 10 in diameter. Male: Unknown. Amblyseius (Proprioseiopsis) tulearensis sp. n. (figs ) Holotype $ (B 27.3) arid 6 paratypes (series 58 and B 27; all 9$) from Corchorus trilocularis (Tiliaceae), This species was found in association with Tetranychus neocaledonicus. botanical garden, Agricultural Station,Tul ar, 2.IV Differential diagnosis. A. tulearensis belongs to Amblyseius (Proprioseiopsis) peltatus Van der Merwe, 1968, comb. n. (figs ) the group of species within the subgenus Proprioseiopsis Muma, 1961, in which only setae J2, and not Zl, are absent, and genu II bears eigth setae. Unfortunately, the latter character has not been considered by a majority of workers, and was never mentioned for American A. asetus (Chant, 1959) (Schuster & Pritchard, 1963; Muma & Amblyseius (Amblyseius) peltatus Van der Merwe, 1968: 119, figs (M 10.10) collected on Urena lobata (Malvaceae) near Ambato Boeni (Majunga province) on 24.IV Denmark, 1970), the descriptions of which fit A. tulearensis rather well. A. asetus (Chant, sensu Schuster, 1966) from the Gal&pagos has genu II with eight setae, but differs from the considerably Diagnosis. specimen agrees with the original description of A. peltatus. This species should be placed in the subgenus Proprioseiopsis original description. Muma, 1961, since the setal J2 pair is absent. Therefore, I prefer to consider A. tulearensis a new species, which resembles A. apheles Van der Merwe, 1968, except Female: Dorsal shield brownish in preparation, for minor differences in the length of some dorsal smooth, 320 long and 230 wide. Length of the setae, and the shape of the spermatheca. Description. Female: Alive brownish red, scutal parts stay yellowish in cleared and mounted specimens. Dorsal shield smooth, not exceptionally sclerotized, 340 long and 230 wide, with at least longer setae: jl 32, j3 61, z4 25, Z4 100, Z5 110, s2 30, s4 100, S2 21. Width of both the genital and ventri-anal shield 102; length of the latter 93. VL1 77 long. Macrosetae on legs: on genu IV 50, tibia IV 36, and basitarsus IV 86, on genu III 32 long. Genu II with seven setae.

22 - BIJDRAGEN TOT DE DIERKUNDE, 46 (1) Figs Amblyseius tulearensis sp. n.: 57, dorsum ; 58, spermatheca ; 59, chelicera ; 60, leg IV ; 61, genital and ventri-anal shield. Figs Amblyseius peltalus Van der Merwe, 1968: 62, leg IV ; 63, sternal shield ; 64, spermatheca.

23 It 102 L. - BLOMMERS PHYTOSEIIDAE FROM MADAGASCAR Remark. should be noted that A. peltatus collected mites could be reared on detached leaf resembles so much A. rosellus comb. n. (Chant, cultures with spider mites as food. 1959) from the Caribbean area, as to be a To this purpose, every sample collected in the synonym of it. Having not seen the type of either, field was split into two parts. A number of I prefer to determine the Malagasy specimen the specimens were preserved immediately for iden- same as the more extensively described African tification. The majority, however, was used for species. rearing experiments. Because it is extremely difficult to recognize different phytoseiids while alive, some (negative) results could have been BIOLOGY lost. It is possible that some species not able to live under the conditions of our rearing methods The main purpose of our study in Madagascar was to search for phytoseiid mites as control agents for noxious tetranychids on local crops. At the time of our arrival, only a single species of was Phytoseiidae described from the island (Chazeau, 1970) and our first goal was to undertake a faunal inventory. This was combined, from the beginning, with trials to determine if the were lost and consequently not preserved. We have made collections in four provinces: Tamatave, Tananarive, and Majunga Tul6ar. The survey in the region of Tamatave concerned especially the spider mites and their predators on citrus and other fruit trees, the results of which have been published elsewhere (Blommers, 1974a; Blommers & Gutierrez, 1975). Fig. 65. Length (individual observations, mean and standard deviation) of the longest macrosetae on leg IV of the female in the Amblyseius rotundus-brevipes group: A. brevipes: A, Tuléar; B, Ambato-Boeni region; C, Tamatave; D, Tananarive. A. rotundus: E, Ambato-Boeni region; F, Tuléar; G, Majunga-Katsepy. (See also table I.)

24 The The - BIJDRAGEN TOT DE DIERKUNDE, 46 (1) Further, we have been concentrating mainly on Tetranychus neocaledonicus Andrd, 1933, as prey species, since it is by far the most important spider mite on the island (Gutierrez, 1974). Tulear Phytoseiid mites collected in the field were offered Tetranychus neocaledonicus as food alone, on detached leaves of either cotton, bean or cassava. This proved to be a quite rigorous way of screening, as only three species survived: Amblyseius masiaka and A. vazimba Blommers & Chazeau, 1974 (cf. Blommers, 1974b) and A. bibens (cf. Blommers & Van Etten, 1975). Other species, notably Amblyseius rotundus, A. brevipes, A. sakalava, Paraphytoseius multidentatus, Phytoseius onilahy and several Typhlodromus species, which were encountered and tested more than once, appeared unable to live with the spider mite as food exclusively. An exception might be Amblyseius tulearensis which found was only once in low numbers, and could be maintained for a few months. Probably, inbreeding was the major reason for its extinction. However, its rareness indicates that this species can only be of minor importance with respect to spider mite regulation. In the same region, another spider mite, Eutetranychus eliei Gutierrez & Helle, 1971, is very common on citrus. No phytoseiids were found in evident association with it. Nearly all our observations and collections concern the centre and immediate environment of the city of Tulear, an area which is irrigated. Phytoseiid mites proved to be extremely the sclerophyllous vegetation rare on on the eastern limestone plateau and in the dunes near the coast, although a wide variety of spider mites occurs there (Gutierrez, pers. comm.). shrub. Several species of phytophagous mites were observed together with them, notably Tetranychus neocaledonicus, Eotetranychus savanae Gutierrez, 1967, Eutetranychus sambiranensis Gutierrez & Helle, 1971, Oligonychus occidentalis Gutierrez, 1969, plus two unknown species the latter genera, Raoiella indica Hirst, unidentified Tarsonemidae. of both 1924 and Amblyseius masiaka, A. tamatavensisand Typhlodromus contiguus were collected simultaneously on adventive sweet potato and bean plants in a patch of degraded dune forest on the road to Mitsinjo, in association with Tetranychus neocaledonicus. western lowlands. A few short visits were made to the cotton growing area near Ambato Boeni and to the outskirts of the deciduous Ankarafantsika forest near Ampijoroa. On all these trips, collected specimens were preserved partly in the field, while the remainder with the plant pieces harbouring them were put into plastic bags and transported to the laboratory in Tananarive. There, the material was sorted and mass rearings were started of as many as possible species by providing the phytoseiid mites on detached bean leaves with not only Tetranychus neocaledonicus but also pollen (Aloe chabaudii: Liliaceae) and beehoney. It appeared possible to rear the following species on this food: Amblyseius rotundus (M 9, M 16), A. brevipes (M 17), A. sundi(m 6), A. masiaka, A. tamatavensis and Typhlodromus contiguus (all M 22), Phytoseius betsiboka (M 30) and Typhlodromus cf. gutierrezichazeaui (M 9, M 16). Amblyseius trichophilus (M 27) and another strain of A. rotundus (Mil) could not be maintainedin this way. After we had established rearings of sufficient size, we experimented to determineif the animals could live with tetranychid mites exclusively, or Majunga with some other food substances. Apart from Amblyseius masiaka, which thrives well with western coastal area. Our survey was spider mites as food (Blommers, 1974b) only one limited to a few days visit to Katsepy, opposite other species was capable to do so: A. sundi. Majunga on the mouth of the Betsiboka River. However, it did much better if honey was The dune vegetation there is more open than near Tulear (cattle grazing), though the precipitation is about five fold (1.5 instead of 0.3 (n annually). Two species were found there: Amblyseius rotundus and Typhlodromus cf. gutierrezi-chazeaui, supplied additionally, like we have observed previously with Amblyseius vazimba (Blommers, 1974b). All other species seemed less disposed to prey on Tetranychus neocaledonicus. If pollen was withheld, the egg production dropped gradually and none or a few juveniles became adult, as is both in rather large numbers on differentsorts of the case for Amblyseius rotundus, A. brevipes, A.

25 - 104 L. BLOMMERS PHYTOSEIIDAE FROM MADAGASCAR was found always roaming solitary in shrubs and trees, including grapevine and but never in figs, the company of spider mites. Nevertheless, it can be reared easily with Tetranychus neocaledonicus tamatavensis and Typhlodromus contiguus. On the contrary, all four species did well if fed with pollen exclusively, although some cannibalism occurred. Phytoseius betsiboka and the Typhlodromus species fed exclusively on the pollen, together with the pollen of Bauhinia sp. (Caesal- even in the presence of spider mites. pinaceae) or Aloe chabaudii as food. Some life We also wanted to ascertain if some of these species would prey on Oligonychus coffeae (Nietner, 1861), a spider which might become mite noxious on some perennial crops (cf. Blommers & Gutierrez, 1975). This spider mite was offered on detached leaves of Ricinus communis, to the exclusion of other food. A clear difference was noted between related Amblyseius rotundus and A. Females of the brevipes. former of which species, both strains were tested, accepted this prey readily and continued egg laying, while the latter species did not eat and succumbed. Juveniles of A. rotundus are annoyed considerably by the webbing of this prey at large densities, but since we have observed them to occur together on frangipani (M 11), a predator/prey relation might exist. Amblyseius masiaka accepts Oligonychus coffeae well as as do A. tamatavensis and prey, Typhlodromus contiguus to a lesser degree. T ananarive history data were determined on this alimentary regime. The duration of the juvenile development (from egg till deutonymph) takes 7.3 days at 23 ± 2 C and 55 ± 5% r.h.; the pre-oviposition period lasts less than three days under the same conditions (J. Chazeau, pers. comm.). The oviposition rate equals about 1.5 eggs/female/day for the first three weeks of the oviposition period at 20± 1 C and 60 ± 10% r.h.; the entire egg-laying period takes more than 50 days, in which a female lays about 60 eggs. Tetranychus neocaledonicus alone is insufficient food to maintain a population of this predacious mite. The juvenile development stops completely and, eventually, the young die. Also, the egg production of the young females drops drastically, in 5 days to 0.5 eggs/day at 25 C. While the observations so far mentioned indicate that Iphiseius degenerans is incapable of controlling Tetranychus neocaledonicus in the field, another spider mite, Oligonychus is coffeae, more susceptible to attack by this predator. This species lives on the same plants from which I. Although Tetranychus neocaledonicus is also common on all sorts of plants, wild or cultivated, in the central highlands of Madagascar, we have never found any phytoseiid there clearly associated with it, excepted Amblyseius bibens (T 25) one time. Rearing experiments, too, never yielded a clear indication to this effect. The following species could be reared on a diet of Tetranychus neocaledonicus and pollen: Phytoseius amba, Iphiseius degenerans, Amblyseius hima (T 1, T 5), A. brevipes (T 8), A. deleoni (T 9) and A. sundi (T 10). Only A. sundi could be maintained on the spider mite alone. Oligonychus coffeae suitable for Phytoseius crinitus(t 19). was not Of the species from this region, Iphiseius degenerans is recorded: notably grapevine, figs, tea etc., and occurs at times in considerable number on commercially treated plantings of the first host in the surroundings of Tananarive. The predacious mite thrives better on O. coffeae without pollen (on leaf arenas of Ricinus communis) than on any other diet tried. Predation is rather elevated: young female predators kill over ten O. coffeae females/day and deposit 2-3 eggs at 22 ± 1 0 C. With Tetranychus neocaledonicus plus pollen, predation drops to less than one female/ day; if pollen is withheld, the predation rises initially to approximately seven prey females in the first 24 hours, but drops thereafter in accordance with the egg production (see above). degenerans was studied in some detail. This Our results agreeroughly with those of Hessein unmistakable, large, wine-red species (1967) on the same predator (from Egypt) with was encountered only at Tananarive. Since it is recorded from several Tetranychus pacificus McGregor, 1919, Oligonychus punicae (Hirst, 1926) and Panonychus citri localities in the Mediterranean (Chant, 1959b) and from high altitude in (McGregor, 1916). However, none of these prey Central Africa (Pritchard & Baker, 1962), one species seem to be optimal food. Addition of might assume that its distribution inside Madagascar is restricted to the central high plateau. It Hymenocyclus pollen always improved the survival of the juveniles.

26 BIJDRAGEN TOT - DE DIERKUNDE, 46 (1) An experiment using two potted Ricinus communis plants in the laboratory showed that Iphiseius degenerans is capable of exterminating Oligonychus coffeae also outside the confinementof a detached leaf culture. II. A taxonomic review of the family Phytoseiidae, with descriptions of 38 new species. Can. Ent., 91 (suppl. 12): CHAZEAU, J., Typhlodromus scytinus n. sp., nouveau phytoseiide de Madagascar (Acariens, Gamasides, Phytoseiidae). Cah. Off. Rech. Sci. Tech. Outre-Mer, Ser. Biol., 12:314. CORPUZ, L. A. & L. RIMANDO, Philippine Amblyseiinae ACKNOWLEDGEMENTS (Phytoseiidae: Acarina). Philipp. Agric., 50: DENMARK, H. A., This study was conducted as a part of project GUA-4 of the Netherlands' University Foundation for International Cooperation (NUFFIC), in collaboration with the ORSTOM centre (Office de la Recherche Scientifique et Technique Outre-Mer) in Tananarive. The author is extremely grateful to Dr. Jean Gutierrez and Mr. Jean Chazeau who have helped him full-heartedly and in every way with his research in Madagascar, and to Mr. Rasoloson Jacques who, with his quiet precision, took care ofthe mass rearings. Revision of the genus Phytoseius Ribaga, 1904 (Acarina: Phytoseiidae). Fla. Dept. Agric. Div. Plant Ind. Bull., 6: EHARA, S., Some predatory mites of the genera Typhlodromus and Amblyseius from Japan (Phytoseiidae). Acarologia, 1: , Phytoseiid mites from Okinawa Island (Acarina Mesostigmata). Mushi, 40: 6782., Some phytoseiid mites from Japan, with description of thirteen new species (Acarina: Mesostigmata). Mushi, 46: EHARA, S. & L. H. Y. LEE, Mites associated with plants in Hong Kong. J. Fac. Educ. Tottori Univ. (Nat. Sci.), 22: REFERENCES EVANS, G. O., The genus Iphiseius Berl. (Acarina ATHIAS-HENRIOT, C., Phytoseiidae et Aceosejidae (Acarina, Gamasina) d'algerie. Bull. Soc. Hist. nat. Afr. N., 48: , Contribution a l'etude des Amblyseius paldarctiques (Acariens Anactinotriches, Phytoseiidae). Bull, scient. Bourgogne, 24: BERLESE, A., Acari, Myriapoda, et Scorpiones hucusque in Italia 54 reperta, (9): [1 3] (Tipografia del seminario, Padova). Bi.OMMERS, L., Five newspecies of phytoseiid mites (Acarina: Mesostigmata) from Southwest Madagascar. Bull, zool. Mus. Univ. Amsterdam, 3: , 1974a. Species of the genus Amblyseius Berlese, 1914, from Tamatave, East Madagascar. Bull. zool. Mus. Univ. Amsterdam, 3: , 1974b. Preliminary studies on two predators (Acarina: Phytoseiidae) ofthe spider mite Tetranychus neocaledonicus Andri (Acarina: Tetranychidae). Z. angew. Ent., 75: BLOMMERS, L. & J. CHAZEAU, Two new species of predator mites of the genus Amblyseius Berlese (Acarina: Phy- Laelaptidae). Proc. zool. Soc. Lond., 124: EVANS, G. O. & D. MACFARLANE, A new mite of the genus Phytoseius Ribaga (Acari: Mesostigmata). Ann. Mag. nat. Hist., (13)4: GUPTA, S. K Three new species of the genus Phytoseius (Acarina: Phytoseiidae) from India. Israel J. agric. Res., 19: GUTIERREZ, J., tude biologique et ecologique de Tetranychus neocaledonicus (Acariens, Tetranychidae): (These de Doctorat d'fitat, Univ. Paris-Sud, France). HESSETN, N. A., Studies on the bionomics of the predacious mite Iphiseius degenerans (Berl.) (Acarina: Phytoseiidae): 1102 (Ph. D. Thesis, Univ. Calif., Riverside, USA). LEON, D. DE, Phytoseiid mites from Puerto Rico with descriptions of new species (Acarina: Mesostigmata). Fla. Ent., 48: LINDQUIST, E. E. & G. O. EVANS, Taxonomic concepts in the Ascidae with a modified setal nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmata). Mem. ent. Soc. Can., 47: 166. toseiidae) from Madagascar. Z. angew. Ent., 75: MERWE, G. G. VAN DER, South African Phytoseiidae (A- BLOMMERS, L. & J. VAN ETTEN, Amblyseius bibens (Acarina: Phytoseiidae), a predator of spider mites (Tetranychidae) in Madagascar. Entomologia exp. appl., 18: carina). I. Nine new species of the genus Amblyseius Berlese. J. ent. Soc. sth. Afr., 28: 5776., A taxonomic study of the family Phytoseiidae (Acari) in South Africa, with contributions to the biology of BLOMMERS, L. & J. GUTIERREZ, Les tetranyques vivant two species. Entomology Mem. Dep. agric. tech. Serv. sur agrumes et avocatiers dans la region de Tamatave repub. S. Afr., 18: [iiv], (Madagascar-est) et quelques-uns de leurs predateurs. Fruits, 30: MUMA, M. H., Phytoseiidae (Acarina) associated with citrus in Florida. Ann. ent. Soc. Am., 48: CHANT, D. A., Descriptions of two new phytoseiid, genera (Acarina: Phytoseiidae), with a note on the genus Phytoseius Ribaga, Can. Ent., 89: a. Description of a new species of Typhlodromus (Acarina: Phytoseiidae), from Eastern Asia. Can. Ent., 91:, Subfamilies, genera and species of Phytoseiidae (Acarina: Mesostigmata). Bull. Fla. State Mus., 5: , Annotated list and keys to Phytoseiidae (Acarina: Mesostigmata) associated with Florida citrus. Fla. agric. Exp. Stn. tech. Bull., 685: , 1959b. Phytoseiid mites (Acarina: Phytoseiidae). Part I., New Phytoseiidae (Acarina: Mesostigmata) from southern Asia. Fla. Ent., 50: Bionomics of seven species of Southeastern England. Part MUMA, M. H. & H. A. DENMARK, Some generic descrip-

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